bioselБиотехнология и селекция растенийPlant Biotechnology and Breeding2658-62662658-6258VIR10.30901/2658-6266-2021-2-o2biosel-133Research ArticleМЕТОДЫ БИОТЕХНОЛОГИИ В СЕМЕНОВОДСТВЕ И СЕЛЕКЦИИ РАСТЕНИЙBIOTECHNOLOGY TECHNIQUES IN SEED PRODUCTION AND PLANT BREEDINGИдентификация генов устойчивости к бурой ржавчине у новых российских сортов мягкой пшеницыIdentification of leaf rust resistance genes in the new Russian varieties of common wheathttps://orcid.org/0000-0001-7948-0307ГультяеваЕ. И.GultyaevaE. I.

196608 Россия, г. Санкт-Петербург, Пушкин, ш. Подбельского, 3

3 Podbelskogo Street, St. Petersburg, Pushkin, 196608 Russia

eigultyaeva@gmail.comhttps://orcid.org/0000-0003-3266-6272ШайдаюкЕ. Л.ShaydayukE. L.

196608 Россия, г. Санкт-Петербург, Пушкин, ш. Подбельского, 3

3 Podbelskogo Street, St. Petersburg, Pushkin, 196608 Russia

Всероссийский научно-исследовательский институт защиты растенийРоссияAll-Russian Institute of Plant ProtectionRussian Federation202107122021421527Copyright © Гультяева Е.И., Шайдаюк Е.Л., 20212021Гультяева Е.И., Шайдаюк Е.Л.Gultyaeva E.I., Shaydayuk E.L.This work is licensed under a Creative Commons Attribution 4.0 License.https://biosel.elpub.ru/jour/article/view/133Актуальность

Актуальность. Бурая ржавчина (возбудитель Puccinia triticina Erikss.) является серьёзным заболеванием пшеницы во всех регионах Российской Федерации. Генетическое разнообразие выращиваемых сортов по типам устойчивости и генам, ее контролирующим, обеспечивает надежную защиту пшеницы от данного патогена. Цель работы – характеристика разнообразия новых российских сортов озимой и яровой мягкой пшеницы по генам устойчивости к бурой ржавчине (Lr-генам).

Материалы и методы

Материалы и методы. Изучены 43 сорта озимой и 25 сортов яровой пшеницы, включенные в Государственный реестр селекционных достижений РФ в 2018-2020-х годах. С использованием молекулярных маркеров нами была проведена идентификация 18 Lr-генов: Lr1, Lr3, Lr9, Lr10, Lr19, Lr20, Lr21, Lr24, Lr25, Lr26, Lr28, Lr29, Lr34, Lr35, Lr37, Lr41 (39), Lr47, Lr66. Фитопатологический тест был использован для уточнения результатов молекулярного анализа.

Результаты

Результаты. У 93% изученных сортов пшеницы выявлены Lr-гены, которые встречались по отдельности или в разных сочетаниях. Это были высоко и частично эффективные гены Lr24, Lr9 и Lr19; гены устойчивости взрослых растений Lr34 и Lr37; малоэффективные гены Lr1, Lr3, Lr10, Lr20 и Lr26. Впервые у сортов российской селекции идентифицирован ген Lr24. Яровой сорт ‘Лидер 80’ с Lr24 рекомендован для возделывания в Западно-Сибирском и Восточно-Сибирском регионах. У высокоустойчивого к бурой ржавчине ярового сорта ‘Силач’ определено эффективное сочетание генов Lr9+Lr26, по отдельности не обеспечивающих устойчивости к патогену. Ген Lr9 выявлен у озимого сорта ‘Герда’, рекомендуемого для возделывания в Северо-Кавказском регионе. Ранее сорта с геном Lr9 на Северном Кавказе не выращивали. Среди сортов пшеницы, проходящих районирование, отмечается увеличение числа образцов, резистентных к бурой ржавчине, защищенных эффективным геном устойчивости взрослых растений Lr37. Гены Lr19, Lr24, Lr26, Lr34, Lr37 сцеплены с эффективными Sr-генами Sr25, Sr24, Sr31, Sr57 и Sr38, что дополнительно обеспечивает новым сортам пшеницы стабильную генетическую защиту от стеблевой ржавчины.

Заключение

Заключение. Полученные сведения представленности Lr-генов в сортах пшеницы следует учитывать в региональных селекционных программах. Своевременная смена генетически защищенных сортов позволяет стабилизировать популяционный состав фитопатогенов и снизить вероятность эпифитотий.

Background

Background. Wheat leaf rust caused by Puccinia triticina Erikss. is a significant wheat disease in all regions of the Russian Federation. The genetic diversity of the cultivated wheat varieties regarding the type of resistance and genes that control it ensures reliable protection of this crop against the pathogen. The aim of this work was to characterize the diversity of new Russian varieties of winter and spring common wheat for leaf rust resistance genes (Lr-genes).

Materials and Methods

Materials and Methods. The research material was represented by 43 varieties of winter and 25 of spring wheat included in the State Register of Selection Achievements of the Russian Federation in 2018-2020.

Results

Results. Using molecular markers, 18 Lr genes were identified: Lr1, Lr3, Lr9, Lr10, Lr19, Lr20, Lr21, Lr24, Lr25, Lr26, Lr28, Lr29, Lr34, Lr35, Lr37, Lr41 (39), Lr47 and Lr66. A phytopathological test was used to clarify the results of molecular analysis. Ninety-three percent of the studied wheat varieties were found to contain Lr genes, either separately or in combinations. These were the highly and partially effective genes Lr24, Lr9, and Lr19, adult plant resistance genes Lr34 and Lr37, and ineffective genes Lr1, Lr3, Lr10, Lr20, and Lr26. The Lr24 gene has been identified for the first time in Russian varieties. The spring variety ‘Leader 80’, harboring this gene, is recommended for cultivation in the West Siberian and East Siberian regions. An effective combination of Lr9 + Lr26 genes, individually overcome by the pathogen, was determined in the spring cultivar ‘Silach’, highly resistant to leaf rust. The Lr9 gene was found in the winter variety ‘Gerda’, which is recommended for cultivation in the North Caucasus region. Previously, the varieties with Lr9 were not grown in the North Caucasus. An increase in the number of leaf rust resistant accessions protected by the effective adult plant resistance gene Lr37 is noted among wheat varieties undergoing regional adaptation testing. Many of the identified Lr genes (Lr19, Lr24, Lr26, Lr34, Lr37) are linked with effective Sr genes (Sr25, Sr24, Sr31, Sr57, and Sr38), which additionally ensures stable genetic protection of wheat against stem rust.

Conclusions

Conclusions. The obtained information about representation of Lr genes in wheat varieties should be used in regional breeding programs. A timely replacement of genetically protected varieties allows stabilizing the populational composition of the phytopathogen and reducing the likelihood of epiphytotics.

молекулярные маркерыустойчивостьLr-геныPuccinia triticinaTriticum aestivummolecular markersresistanceLr genesPuccinia triticinaTriticum aestivumРабота выполнена в рамках государственного задания ВИЗР, проект 0665-2019-0015 «Грибы – патогены экономически значимых растений в России: разнообразие, методы идентификации и мониторинга, взаимоотношения с растениями-хозяевами».The research was performed within the framework of the State Assignment to VIZR, Project No. 0665-2019-0015 “Fungal pathogens of economically significant plants in Russia: diversity, methods of identification and monitoring, relationships with host plantsˮ.ReferencesАгапова В.Д., Ваганова О.Ф., Волкова Г.В. Эффективность ювенильных генов устойчивости к возбудителю бурой ржавчины озимой пшеницы в фазу проростков в условиях юга России. Международный научно-исследовательский журнал. 2020;8(98):163-167 DOI: 10.23670/IRJ.2020.98.8.023Agapova V.D., Vaganova O.F., Volkova G.V. The efficiency of juvenile genes of orange leaf rust resistance of winter wheat during the germinal phase in the climate of the Russian South. International Research Journal. 2020;8(98):163-167. [in Russian] DOI: 10.23670/IRJ.2020.98.8.023Atia M.A.M., El-Khateeb E.A., Abd El-Maksoud R.M., Abou-Zeid M.A., Salah A., Abdel-Hamid A.M.E. Mining of leaf rust resistance genes content in Egyptian bread wheat collection. Plants. 2021;10(7);1378. DOI: 10.3390/plants10071378Atia M.A.M., El-Khateeb E.A., Abd El-Maksoud R.M., Abou-Zeid M.A., Salah A., Abdel-Hamid A.M.E. Mining of leaf rust resistance genes content in Egyptian bread wheat collection. Plants. 2021;10(7);1378. DOI: 10.3390/plants10071378Brown-Guedira G., Sing S. Leaf Rust Resistance Gene Lr39. MasWheat. Marker Assisted Selection in Wheat. 2019. Available from: https://maswheat.ucdavis.edu/protocols/Lr39 [accessed on 10 May 2021].Brown-Guedira G., Sing S. Leaf Rust Resistance Gene Lr39. MasWheat. Marker Assisted Selection in Wheat. 2019. Available from: https://maswheat.ucdavis.edu/protocols/Lr39 [accessed on 10 May 2021].Chelkowski J., Golka L., Stepien L. Application of STS markers for leaf rust resistance genes in near-isogenic lines of spring wheat cv. Thatcher. Journal of Applied Genetics. 2003;44:323-338.Chelkowski J., Golka L., Stepien L. Application of STS markers for leaf rust resistance genes in near-isogenic lines of spring wheat cv. Thatcher. Journal of Applied Genetics. 2003;44:323-338.Cherukuri D.P., Gupta S.K., Charpe A., Koul S., Prabhu K.V., Singh R.B., Haq Q.M.R. Molecular mapping of Aegilops speltoides derived leaf rust resistance gene Lr28 in wheat. Euphytica. 2005;143:19-26. DOI: 10.1007/s10681-005-1680-6Cherukuri D.P., Gupta S.K., Charpe A., Koul S., Prabhu K.V., Singh R.B., Haq Q.M.R. Molecular mapping of Aegilops speltoides derived leaf rust resistance gene Lr28 in wheat. Euphytica. 2005;143:19-26. DOI: 10.1007/s10681-005-1680-6Dakouri A., McCallum B.D., Radovanovic N., Cloutier S. Molecular and phenotypic characterization of seedling and adult plant leaf rust resistance in a world wheat collection. Molecular Breeding. 2013;32:663-677. DOI: 10.1007/s11032-013-9899-8Dakouri A., McCallum B.D., Radovanovic N., Cloutier S. Molecular and phenotypic characterization of seedling and adult plant leaf rust resistance in a world wheat collection. Molecular Breeding. 2013;32:663-677. DOI: 10.1007/s11032-013-9899-8Дорохов Д.Б., Клоке Э. Быстрая и экономичная технология RAPD анализа растительных геномов. Генетика. 33(4):443-450.Dorokhov D.B., Kloke E. A rapid and economical technology of RAPD analysis of plant genomes. Russian Journal of Genetics. 1997;33(4):358-365. [in Russian]Дьяков Ю.Т. Популяционная биология фитопатогенных грибов. Москва: Издательский дом «Муравей»; 1998.Dyakov Yu.T. Population biology of phytopathogenic fungi (Populyatsionnaya biologiya fitopatogennykh gribov). Moscow: “Muraveyˮ Publishing House; 1998. [in Russian]Fritz A. Leaf Rust Resistance Gene Lr21. Marker Assisted Selection in Wheat (MasWheat). 2019. Available from: http://maswheat.ucdavis.edu/protocols/Lr21 [accessed on 10 May 2021].Fritz A. Leaf Rust Resistance Gene Lr21. Marker Assisted Selection in Wheat (MasWheat). 2019. Available from: http://maswheat.ucdavis.edu/protocols/Lr21 [accessed on 10 May 2021].Gregáňová Ž., Kraic J., Gálová Z. Diagnostic of wheat leaf rust resistance genes by DNA markers and their application in MAS. Czech Journal of Genetics and Plant Breeding. 2003;39:127-129. DOI: 10.17221/3730-CJGPBGregáňová Ž., Kraic J., Gálová Z. Diagnostic of wheat leaf rust resistance genes by DNA markers and their application in MAS. Czech Journal of Genetics and Plant Breeding. 2003;39:127-129. DOI: 10.17221/3730-CJGPBГультяева Е.И. Методы идентификации генов устойчивости пшеницы к бурой ржавчине с использованием ДНК-маркеров и характеристика эффективности Lr-генов. Санкт-Петербург; 2012.Gultyaeva E.I. Methods for identification of wheat leaf rust resistance genes using DNA markers and characterization of the efficiency of Lr genes (Metody identifikatsii genov ustoychivosti pshenitsy k buroy rzhavchine s ispolzovaniyem DNK-markerov i kharakteristiki effektivnosti Lr-genov). St. Petersburg; 2012. [in Russian]Гультяева Е.И., Баранова О.А., Дмитриев А.П. Вирулентность и структура популяций Puccinia triticina в Российской Федерации в 2007 году. Вестник защиты растений. 2009;4:33-38.Gultyaeva E.I., Baranova О.А., Dmitriev А.P.V Virulence and population structure of Puccinia triticina in Russian Federation in 2007. Plant Protection News. 2009;4:33-38. [in Russian]Гультяева Е.И., Садовая А.С. Селекция на устойчивость к бурой ржавчине в России. Защита и карантин растений. 2014;10:24-26.Gultyaeva E.I., Sadovaya A.S. Breeding of common wheat for resistance to brown rust of wheat in Russia. Plant Protection and Quarantine. 2014;10:24-26. [in Russian]Gultyaeva E.I., Shaydayuk E.L., Gannibal Ph.B. Leaf rust resistance genes in wheat cultivars registered in Russia and their influence on adaptation processes in pathogen populations. Agriculture. 2021;11(4):319. DOI: 10.3390/agriculture11040319Gultyaeva E.I., Shaydayuk E.L., Gannibal Ph.B. Leaf rust resistance genes in wheat cultivars registered in Russia and their influence on adaptation processes in pathogen populations. Agriculture. 2021;11(4):319. DOI: 10.3390/agriculture11040319Gultyaeva E.I., Shaydayuk E.L., Kosman E.G. Regional and temporal differentiation of virulence phenotypes of Puccinia triticina from common wheat in Russia during the period 2001-2018. Plant Pathology. 2020;69(5):860-871. DOI: 10.1111/ppa.13174Gultyaeva E.I., Shaydayuk E.L., Kosman E.G. Regional and temporal differentiation of virulence phenotypes of Puccinia triticina from common wheat in Russia during the period 2001-2018. Plant Pathology. 2020;69(5):860-871. DOI: 10.1111/ppa.13174Gupta S.K., Charpe A., Koul S., Prabhu K.V., Haq Q.M.R. Affiliations expand development and validation of molecular markers linked to an Aegilops umbellulata-derived leaf-rust-resistance gene, Lr9, for marker-assisted selection in bread wheat. Genome. 2005;48(5):823-830. DOI: 10.1139/g05-051Gupta S.K., Charpe A., Koul S., Prabhu K.V., Haq Q.M.R. Affiliations expand development and validation of molecular markers linked to an Aegilops umbellulata-derived leaf-rust-resistance gene, Lr9, for marker-assisted selection in bread wheat. Genome. 2005;48(5):823-830. DOI: 10.1139/g05-051Gupta S.K., Charpe A., Prabhu K.W., Haque O.M.R. Identification and validation of molecular markers linked to the leaf rust resistance gene Lr19 in wheat. Theoretical and Applied Genetics. 2006;113(6):1027-1036. DOI: 10.1007/s00122-006-0362-7Gupta S.K., Charpe A., Prabhu K.W., Haque O.M.R. Identification and validation of molecular markers linked to the leaf rust resistance gene Lr19 in wheat. Theoretical and Applied Genetics. 2006;113(6):1027-1036. DOI: 10.1007/s00122-006-0362-7Halpin C. Gene stacking in transgenic plants – the challenge for 21st century plant biotechnology. Plant Biotechnology Journal. 2005;3:141-155. doi: 10.1111/j.1467-7652.2004.00113.xHalpin C. Gene stacking in transgenic plants – the challenge for 21st century plant biotechnology. Plant Biotechnology Journal. 2005;3:141-155. doi: 10.1111/j.1467-7652.2004.00113.xHelguera M., Khan I.A., Dubcovsky J. Development of PCR markers for wheat leaf rust resistance gene Lr47. Theoretical and Applied Genetics. 2000;101: 625-631. DOI: 10.1007/s001220051524Helguera M., Khan I.A., Dubcovsky J. Development of PCR markers for wheat leaf rust resistance gene Lr47. Theoretical and Applied Genetics. 2000;101: 625-631. DOI: 10.1007/s001220051524Helguera M., Khan I.A., Kolmer J., Lijavetzky D., Zhong-qi L., Dubcovsky J. PCR assays for the Lr37–Yr17–Sr38 cluster of rust resistance genes and their use to develop isogenic hard red spring wheat lines. Crop Science. 2003;43(5):1839-1847. DOI: 10.2135/cropsci2003.1839Helguera M., Khan I.A., Kolmer J., Lijavetzky D., Zhong-qi L., Dubcovsky J. PCR assays for the Lr37–Yr17–Sr38 cluster of rust resistance genes and their use to develop isogenic hard red spring wheat lines. Crop Science. 2003;43(5):1839-1847. DOI: 10.2135/cropsci2003.1839Herrera-Foessel S., Singh R.P., Huerta-Espino J., William M., Rosewarne G., Djurle A., Yuen J. Identification and mapping of Lr3 and a linked leaf rust resistance gene in durum wheat. Crop Science. 2007;47(4):1459-1466. DOI: 10.2135/cropsci2006.10.0663Herrera-Foessel S., Singh R.P., Huerta-Espino J., William M., Rosewarne G., Djurle A., Yuen J. Identification and mapping of Lr3 and a linked leaf rust resistance gene in durum wheat. Crop Science. 2007;47(4):1459-1466. DOI: 10.2135/cropsci2006.10.0663Lagudah E.S., McFadden H., Singh R.P., Huerta-Espino J. Bariana H.S., Spielmeyer W. Molecular genetic characterization of the Lr34/Yr18 slow rusting resistance gene region in wheat. Theoretical and Applied Genetics. 2006;114:21-30. DOI: 10.1007/s00122-006-0406-zLagudah E.S., McFadden H., Singh R.P., Huerta-Espino J. Bariana H.S., Spielmeyer W. Molecular genetic characterization of the Lr34/Yr18 slow rusting resistance gene region in wheat. Theoretical and Applied Genetics. 2006;114:21-30. DOI: 10.1007/s00122-006-0406-zMago R., Bariana H.S., Dundas I.S. Development or PCR markers for the selection of wheat stem rust resistance genes Sr24 and Sr26 in diverse wheat germplasm. Theoretical and Applied Genetics. 2005;111(3):496-504. DOI: 10.1007/s00122-005-2039-zMago R., Bariana H.S., Dundas I.S. Development or PCR markers for the selection of wheat stem rust resistance genes Sr24 and Sr26 in diverse wheat germplasm. Theoretical and Applied Genetics. 2005;111(3):496-504. DOI: 10.1007/s00122-005-2039-zMago R., Zhang P., Bariana H.S., Verlin U.K., Ellis J.G., Dundas I.S. Development of wheat lines carrying stem rust resistance gene Sr39 with reduced Aegilops speltoides chromatin and simple PCR markers for marker-assisted selection. Theoretical and Applied Genetics. 2009;119(8):1441-50. DOI: 10.1007/s00122-009-1146-7.Mago R., Zhang P., Bariana H.S., Verlin U.K., Ellis J.G., Dundas I.S. Development of wheat lines carrying stem rust resistance gene Sr39 with reduced Aegilops speltoides chromatin and simple PCR markers for marker-assisted selection. Theoretical and Applied Genetics. 2009;119(8):1441-50. DOI: 10.1007/s00122-009-1146-7.Marais G.F., Bekker T.A., Eksteen A., McCallum B., Fetch T., Marais A.S. Attempts to remove gametocidal genes co-transferred to common wheat with rust resistance from Aegilops speltoides. Euphytica 2010;171(1):71-85. DOI: 10.1007/s10681-009-9996-2Marais G.F., Bekker T.A., Eksteen A., McCallum B., Fetch T., Marais A.S. Attempts to remove gametocidal genes co-transferred to common wheat with rust resistance from Aegilops speltoides. Euphytica 2010;171(1):71-85. DOI: 10.1007/s10681-009-9996-2McCallum B.D., Hiebert C.W., Cloutier S., Bakkeren G., Rosa S.B., Humphreys D.G., Marais J.F., McCartney C.A., Panwar V., Rampitsch C., Saville B.J., Wang X. A review of wheat leaf rust research and the development of resistant cultivars in Canada. Canadian Journal of Plant Pathology. 2016;38(1):1-18. DOI: 10.1080/07060661.2016.1145598McCallum B.D., Hiebert C.W., Cloutier S., Bakkeren G., Rosa S.B., Humphreys D.G., Marais J.F., McCartney C.A., Panwar V., Rampitsch C., Saville B.J., Wang X. A review of wheat leaf rust research and the development of resistant cultivars in Canada. Canadian Journal of Plant Pathology. 2016;38(1):1-18. DOI: 10.1080/07060661.2016.1145598McIntosh R.A., Dubcovsky J., Rogers W.J., Xia X.C., Raupp W.J. Сatalogue of gene symbols for wheat: 2020 supplement. Annual Wheat Newsletter. 2020;66:109-128. Available from: https://wheat.pw.usda.gov/ggpages/awn/66/AWNVOL66.pdf [accessed on 11 May 2021].McIntosh R.A., Dubcovsky J., Rogers W.J., Xia X.C., Raupp W.J. Сatalogue of gene symbols for wheat: 2020 supplement. Annual Wheat Newsletter. 2020;66:109-128. Available from: https://wheat.pw.usda.gov/ggpages/awn/66/AWNVOL66.pdf [accessed on 11 May 2021].McIntosh R.A., Wellings C.R., Park R.F. Wheat rusts: An atlas of resistance genes. Australia: CSIRO; 1995. DOI: 10.1071/9780643101463McIntosh R.A., Wellings C.R., Park R.F. Wheat rusts: An atlas of resistance genes. Australia: CSIRO; 1995. DOI: 10.1071/9780643101463Мешкова Л.В., Россеева Л.П., Коренюк Е.А., Белан И.А. Динамика распространения патотипа возбудителя бурой ржавчины, вирулентного к сортам пшеницы с геном Lr9 в Омской области. Микология и фитопатология. 2012;46:397-400.Meshkova L.V., Rosseeva L.P., Korenyuk E.A., Belan I.A. Dynamics of distribution of the wheat leaf rust pathotypes virulent to the cultivars with Lr9 gene in Omsk region. Mycology and Phytopathology. 2012;46:397-400. [in Russian]Morgounov A., Ablova L., Babayants O., Babayants L., Bespalova L., Khudokormova Zh., Litvinenko N., Shamanin V., Syukov V. Genetic protection of wheat rusts and development of resistant varieties in Russia and Ukraine. Euphytica. 2011;179(2):297-311. DOI: 10.1007/s10681-010-0326-5Morgounov A., Ablova L., Babayants O., Babayants L., Bespalova L., Khudokormova Zh., Litvinenko N., Shamanin V., Syukov V. Genetic protection of wheat rusts and development of resistant varieties in Russia and Ukraine. Euphytica. 2011;179(2):297-311. DOI: 10.1007/s10681-010-0326-5Neu C., Stein N., Keller B. Genetic mapping of the Lr20-Pm1 resistance locus reveals suppressed recombination on chromosome arm 7AL in hexaploid wheat. Genome. 2002;45(4):737-744. DOI: 10.1139/g02-040Neu C., Stein N., Keller B. Genetic mapping of the Lr20-Pm1 resistance locus reveals suppressed recombination on chromosome arm 7AL in hexaploid wheat. Genome. 2002;45(4):737-744. DOI: 10.1139/g02-040Pestsova E., Ganal M.W., Röder M.S. Isolation and mapping of microsatellite markers specific for the D genome of bread wheat. Genome. 2000;43(4):689-697. DOI: 10.1139/g00-042Pestsova E., Ganal M.W., Röder M.S. Isolation and mapping of microsatellite markers specific for the D genome of bread wheat. Genome. 2000;43(4):689-697. DOI: 10.1139/g00-042Плотникова Л.Я., Штубей Т.Ю. Эффективность генов возрастной устойчивости пшеницы к бурой ржавчине Lr22b, Lr34, Lr37 в Западной Сибири и цитофизиологическая основа их действия. Вавиловский журнал генетики и селекции. 2012;16(1):123-131. DOI: 10.1134/S2079059713010115.Plotnikova L.Ya., Shtubey T.Yu. Effectiveness of the wheat Lr22b, Lr34, and Lr37 genes for adult plant resistance to leaf rust in west Siberia and the cytophysiological basis of their action. Vavilov Journal of Genetics and Breeding. 2012;16(1):123-131. [in Russian] DOI: 10.1134/S2079059713010115Procunier J.D., Townley-Smith T.F., Fox S., Prashar S., Gray M., Kim W.K., Czarnecki E., Dyck P.L. PCR-based RAPD/DGGE markers linked to leaf rust resistance genes Lr29 and Lr25 in wheat (Triticum aestivum L.). Journal of Genetics and Breeding. 1995;49:87-92.Procunier J.D., Townley-Smith T.F., Fox S., Prashar S., Gray M., Kim W.K., Czarnecki E., Dyck P.L. PCR-based RAPD/DGGE markers linked to leaf rust resistance genes Lr29 and Lr25 in wheat (Triticum aestivum L.). Journal of Genetics and Breeding. 1995;49:87-92.Qiu J.W., Schürch A.C., Yahiaoui N., Dong L.L., Fan H.J., Zhang Z.J., Keller B., Ling H.Q. Physical mapping and identification of a candidate for the leaf rust resistance gene Lr1 of wheat. Theoretical and Applied Genetics. 2007;115:159-168. DOI: 10.1007/s00122-007-0551-zQiu J.W., Schürch A.C., Yahiaoui N., Dong L.L., Fan H.J., Zhang Z.J., Keller B., Ling H.Q. Physical mapping and identification of a candidate for the leaf rust resistance gene Lr1 of wheat. Theoretical and Applied Genetics. 2007;115:159-168. DOI: 10.1007/s00122-007-0551-zСанин С.С. Эпифитотии болезней зерновых культур: теория и практика: избранные труды. Москва, 2012.Sanin S.S. Epiphytotics of cereal crop diseases: theory and practice: selected works (Epifitotii boleznej zernovyh kultur: teoriya i praktika: izbrannye trudy). Moscow; 2012. [in Russian]Schachermayr G., Feuillet C., Keller B. Molecular markers for the detection of the wheat leaf rust resistance gene Lr10 in diverse genetic backgrounds. Molecular Breeding. 1997;3:65-74.Schachermayr G., Feuillet C., Keller B. Molecular markers for the detection of the wheat leaf rust resistance gene Lr10 in diverse genetic backgrounds. Molecular Breeding. 1997;3:65-74.Serfling A., Krämer I., Lind V., Schliephake E., Ordon F. Diagnostic value of molecular markers for Lr genes and characterization of leaf rust resistance of German winter wheat cultivars with regard to the stability of vertical resistance. European Journal of Plant Pathology. 2011;130:559-575. DOI: 10.1007/s10658-011-9778-2Serfling A., Krämer I., Lind V., Schliephake E., Ordon F. Diagnostic value of molecular markers for Lr genes and characterization of leaf rust resistance of German winter wheat cultivars with regard to the stability of vertical resistance. European Journal of Plant Pathology. 2011;130:559-575. DOI: 10.1007/s10658-011-9778-2Servin B., Martin O.C.; Mézard M., Hospital F. Toward a theory of marker-assisted gene pyramiding. Genetics. 2004:68(1):513-523. doi: 10.1534/genetics.103.023358.Servin B., Martin O.C.; Mézard M., Hospital F. Toward a theory of marker-assisted gene pyramiding. Genetics. 2004:68(1):513-523. doi: 10.1534/genetics.103.023358.Seyfarth R., Feuillt C., Shachermayr G., Messmer M., Winzeler M., Keller B. Molecular mapping of the adult plant leaf rust resistance gene Lr13 in wheat (Triticum aestivum L.). Journal of Genetics and Breeding. 2000;54(3):193-198.Seyfarth R., Feuillt C., Shachermayr G., Messmer M., Winzeler M., Keller B. Molecular mapping of the adult plant leaf rust resistance gene Lr13 in wheat (Triticum aestivum L.). Journal of Genetics and Breeding. 2000;54(3):193-198.Сибикеев С.Н., Бадаева Е.Д., Гультяева Е.И., Дружин А.Е., Шишкина А.А., Драгович А.Ю., Крупин П.Ю., Карлов Г.И., Кхуат Т.М., Дивашук М.Г. Cравнительный анализ 6Аgi и 6Аgi2 хромосом Аgropyron intermedium (Host) Beauv у сортов и линий мягкой пшеницы с пшенично-пырейными замещениями. Генетика. 2017;53(3):98-309. DOI: 10.7868/S0016675817030110Sibikeev S.N., Druzhin A.E., Badaeva E.D., Shishkina A.A., Dragovich A.Y., Gultyaeva E.I., Kroupin P.Y., Karlov G.I., Khuat T.M., Divashuk M.G. Comparative analysis of Agropyron intermedium (Host) Beauv 6Agi and 6Agi2 chromosomes in bread wheat cultivars and lines with wheat–wheatgrass substitutions. Russian Journal of Genetics. 2017;53(3):314-324. [in Russian] DOI: 10.7868/S0016675817030110Сибикеев С.Н., Крупнов В.А. Эволюция листовой ржавчины и защита от нее пшеницы в Поволжье. Вестник Саратовского госагроуниверситета им. Вавилова. 2007;(Спецвыпуск):92-94.Sibikeev S.N., Krupnov V.A. Evolution of leaf rust and protection from it in the Volga region. The Bulletin of Saratov State Agrarian University in honor of N.I. Vavilov. 2007;(Special edition):92-94. [in Russian]Сибикеев С.Н., Маркелова Т.С., Дружин А.Е., Веденеева М.Л., Сингх Д. Оценка набора интрогрессивных линий яровой мягкой пшеницы селекции НИИСХ Юго-Востока на устойчивость к расе стеблевой ржавчине Ug99+Sr24 (TTKST). Доклады РАСХН. 2011;(2):3-5.Sibikeev S.N., Markelova T.S., Druzhin A.E., Vedeneeva M.L., Singkh D. Evaluation of a set of introgressive spring bread wheat lines developed for resistance to stem rust race ug99 + Sr24 (TTKST) at the Agricultural Research Institute for the Southeast Region. Russian Agricultural Sciences. 2011;37(2):95-97. [in Russian]Singh S.K., Sahoo J.P., Swain E. A review on gene stacking in crop plant. Journal of Pharmacognosy and Phytochemistry. 2018;7(4):1862-1865.Singh S.K., Sahoo J.P., Swain E. A review on gene stacking in crop plant. Journal of Pharmacognosy and Phytochemistry. 2018;7(4):1862-1865.Singh R.P., Trethowan R. Breeding spring bread wheat for irrigated and rainfed production systems of developing world. In: Kang M., Priyadarshan P.M. (eds). Breeding major food staples. Ames (Iowa): Blackwell; 2007. p.109-140.Singh R.P., Trethowan R. Breeding spring bread wheat for irrigated and rainfed production systems of developing world. In: Kang M., Priyadarshan P.M. (eds). Breeding major food staples. Ames (Iowa): Blackwell; 2007. p.109-140.Сочалова Л.П., Лихенко И.Е. Оценка устойчивости к бурой ржавчине изогенных по генам Lr-линий и сортов пшеницы в условиях новосибирской области. Достижения науки и техники АПК. 2016;30(3):46-50.Sochalova L.P., Likhenko I.E. Evaluation of resistance to brown rust of Lr-lines and varieties of wheat, isogenic in genes, under conditions of Novosibirsk region. Dostizheniya nauki i tekhniki APK = Achievements of science and technology of AIC. 2016;30(3):46-50. [in Russian]Государственный реестр селекционных достижений, допущенных к использованию. Т.1. «Сорта растений» (официальное издание). Москва: ФГБНУ «Росинформагротех»; 2021. URL : https://gossortrf.ru/gosreestr/ [дата обращения: 10.05.2021]State Register for Selection Achievements Admitted for Usage (National List). Vol.1 "Plant varieties" (official publication). Moscow: FGBNU "Rosinformagrotekh"; 2021. [in Russian] URL : https://gossortrf.ru/gosreestr/ [дата обращения: 10.05.2021]Taverniers I., Papazova N., Bertheau Y., De Loose M., Holst-Jensen A. Gene stacking in transgenic plants: towards compliance between definitions, terminology, and detection within the EU regulatory framework. Environmental Biosafety Research. EDP Sciences. 2008;7(4):197-218. DOI: 10.1051/ebr:2008018Taverniers I., Papazova N., Bertheau Y., De Loose M., Holst-Jensen A. Gene stacking in transgenic plants: towards compliance between definitions, terminology, and detection within the EU regulatory framework. Environmental Biosafety Research. EDP Sciences. 2008;7(4):197-218. DOI: 10.1051/ebr:2008018Tomkowiak A., Skowrońska R., Buda A., Kurasiak-Popowska D., Nawracała J., Kowalczewski L., Pluta M., Radzikowska D. Identification of leaf rust resistance genes in selected wheat cultivars and development of multiplex PCR. Open Life Sciences. 2019;14(1):327-334. DOI: 10.1515/biol-2019-0036Tomkowiak A., Skowrońska R., Buda A., Kurasiak-Popowska D., Nawracała J., Kowalczewski L., Pluta M., Radzikowska D. Identification of leaf rust resistance genes in selected wheat cultivars and development of multiplex PCR. Open Life Sciences. 2019;14(1):327-334. DOI: 10.1515/biol-2019-0036Тырышкин Л.Г., Курбанова П.М. Возможность использования молекулярных маркеров F1.2245, GWM630, XGWM130 и XBARC352 для идентификации генов устойчивости мягкой пшеницы к листовой ржавчине. Известия Санкт-Петербургского государственного аграрного университета. 2009;15:34-38.Tyryshkin L.G., Kurbanova P.M. Possibility of using molecular markers F1.2245, GWM630, XGWM130 and XBARC352 to identify genes for resistance of common wheat to leaf rust (Vozmozhnost ispolzovaniya molekulyarnykh markerov F1.2245, GWM630, XGWM130 I XBARC352 dlya identifikatsii genov ustoychivosti myagkoy pshenitsy k listovoy rzhavchine). Izvestiya Sankt-Peterburgskogo Gosudarstvennogo Agrarnogo Universiteta = Bulletin of the St. Petersburg State Agrarian University. 2009;15:34-38. [in Russian]Vida G., Gál M., Uhrin A., Veisz O., Hasan Syed N., Flavell A.J., Wang Z., Bedò Z. Molecular markers for the identification of resistance genes and marker-assisted selection in breeding wheat for leaf rust resistance. Euphytica. 2009;170(1-2):67-76. DOI: 10.1007/s10681-009-9945-0Vida G., Gál M., Uhrin A., Veisz O., Hasan Syed N., Flavell A.J., Wang Z., Bedò Z. Molecular markers for the identification of resistance genes and marker-assisted selection in breeding wheat for leaf rust resistance. Euphytica. 2009;170(1-2):67-76. DOI: 10.1007/s10681-009-9945-0Weng Y., Azhaguvel P., Devkota R.N., Rudd J.C. PCR based markers for detection of different sources of 1AL.1RS and 1BL.1RS wheat-rye translocations in wheat background. Plant Breeding. 2007;126(5):482-486. DOI: 10.1111/j.1439-0523.2007.01331.xWeng Y., Azhaguvel P., Devkota R.N., Rudd J.C. PCR based markers for detection of different sources of 1AL.1RS and 1BL.1RS wheat-rye translocations in wheat background. Plant Breeding. 2007;126(5):482-486. DOI: 10.1111/j.1439-0523.2007.01331.xZhemchuzhina A., Kurkova N. Structure of population of Puccinia triticina in various regions of Russia in 2006-2008. 8th International Wheat Conference: Abstracts of oral and poster presentations; 2010 June 01-04; St. Petersburg, Russia. St. Petersburg: VIR; 2010. p.279.Zhemchuzhina A., Kurkova N. Structure of population of Puccinia triticina in various regions of Russia in 2006-2008. 8th International Wheat Conference: Abstracts of oral and poster presentations; 2010 June 01-04; St. Petersburg, Russia. St. Petersburg: VIR; 2010. p.279.

The authors declare that there are no conflicts of interest present.